1 Shifts in Sapling Regeneration Over 25 Years in Forest Ecosystems Of

Shifts in sapling regeneration over 25 years in forest ecosystems of Appalachian Ohio

THESIS

Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in

the Graduate School of The Ohio State University

Savannah L. Ballweg

Graduate Program in Environment and Natural Resources

The Ohio State University

2020

Master’s Examination Committee

Dr. David M. Hix, Advisor

Dr. Stephen N. Matthews

Dr. Roger A. Williams

Copyrighted by

Savannah L. Ballweg

2020

Abstract

Many Quercus-Carya forests in the eastern United States are experiencing compositional shifts due to the failure of the overstory species to regenerate, while mesophytic species, i.e., Acer rubrum, Acer saccharum, and Fagus grandifolia, increasingly dominate their regeneration layers. The Wayne National Forest (WNF) of southeastern Ohio is largely

Quercus-Carya forest ecosystem types, although the sapling regeneration is often dominated by the aforementioned mesophytic species. Through the 2018 resampling of

98 permanent plots established on the Marietta Unit of the WNF in 1993, this research investigates changes in sapling species composition and abundance. Statistically significant changes were observed in seven of the species in the total sapling layer: Acer rubrum, Acer saccharum, Aesculus flava, Carya glabra, Cornus florida, Fagus grandifolia, and Hamamelis virginiana. Within the subcategory of small saplings (stems

0.1-5.0 cm DBH), the ten species that had statistically significant changes were Acer rubrum, Acer saccharum, Carpinus caroliniana, Carya glabra, Cornus florida, Fagus grandifolia, Hamamelis virginiana, Nyssa sylvatica, Prunus serotina, and Ulmus rubra.

In the subcategory of large saplings (stems 5.0-10.0 cm DBH), there were two species that had statistically significant changes: Cornus florida and Fagus grandifolia. Overall, the mesophytic species Acer rubrum, Acer saccharum, and Fagus grandifolia continue to dominate the sapling layers of the sampled forests. Fagus grandifolia had significant ii differences in total saplings, as well as both small and large sapling subcategories. In all three classes, Fagus grandifolia increased in stems per hectare and relative density. In the total sapling and small sapling categories, Fagus grandifolia became the most abundant species. Acer rubrum and Acer saccharum had statistically significant changes in total and small sapling categories, where the stems per hectare and relative densities of both species decreased. Acer saccharum and Acer rubrum respectively remained the first and second most abundant in the large sapling subcategory. Comparatively, none of our sparse Quercus species had significant differences in abundance between sampling periods, and Carya glabra had significant changes in total and in small sapling size- classes, with decreasing stems per hectare and relative densities. Our research sought to quantitatively evaluate shifts in sapling composition and abundance between two sampling periods, and found continued prominence of mesophytic species, including the increasing abundance of Fagus grandifolia.

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Acknowledgments

I am very thankful for the support of my advisor, Dr. David Hix, for his constant guidance with my research and education. Being able to continue the research project he played such a large role in is an honor. Dr. Stephen Matthews has also been a fundamental part of my research experience, and has always been eager to lend a hand. I cannot imagine how different my work on the Wayne National Forest would have been without their assistance and counsel. I also owe a special thanks to Dr. Roger Williams, for joining Dr. Hix and Dr. Matthews on my thesis committee. I am very appreciative of the opportunity to have his expertise and perspective applied to this project.

I would also like to express thanks to the field technicians who made this research possible. Hayley Bukala, Alec Dumbauld, Isaac Knowles, and Chas Parise were absolutely essential over the two years I spent with this project. I am also grateful for the help of my predecessor Don Radcliffe, and for the work of Jim Palus and Erin Andrew before him. I would also like to thank my undergraduate advisor, Dr. Carolyn Keiffer, for her continued encouragement of my academic endeavors over the years.

Vita

2018……………………………. B.A. Botany, Environmental Science, College of Arts and Science, Miami University: Oxford, Ohio 2018-2020……………………...Graduate Teaching Associate, School of Environment and Natural Resources, The Ohio State University: Columbus, Ohio

Field of Study

Major Field: Environment and Natural Resources

Specialization: Forest Science

Table of Contents Abstract ...... ii Acknowledgments ...... iv Vita ...... v List of Tables ...... vii List of Figures ...... viii Chapter 1. Introduction ...... 1 1.1 Background ...... 1 1.2 Questions and Hypotheses ...... 3 Chapter 2. Study Area ...... 5 2.1 Climate ...... 5 2.3 Geology and Soils ...... 6 2.4 Vegetation ...... 6 Chapter 3. Methods ...... 8 3.1 Study Design ...... 8 3.2 Sampling Protocol ...... 9 3.3 Data Analyses ...... 11 Chapter 4. Results ...... 13 Chapter 5. Discussion ...... 17 Bibliography ...... 34 Appendix: Sapling species abundance during two sampling periods ...... 43

List of Tables

Table 1. Species of sapling size sampled during the two time periods (1993 and 2018) on the Marietta Unit of the Wayne National Forest by Latin name and species code ...... 21

Table 2. Wilcoxon signed-rank test results for all saplings (both size classes combined) 22

Table 3. Wilcoxon signed-rank test results for small saplings ...... 23

Table 4. Wilcoxon signed-rank test results for large saplings ...... 24

Table 5. Sapling (both size classes combined) stems per hectare (SPH) by year of sampling ...... 25

Table 6. Relative density % (RD) of species for all saplings (both size classes combined)

...... 26

Table 7. Small saplings per hectare (SPH) by year of sampling ...... 27

Table 8. Relative density % (RD) of species for small saplings by year of sampling ...... 28

Table 9. Large saplings per hectare (SPH) by year of sampling ...... 29

Table 10. Relative density % (RD) of species for large saplings by year of sampling ..... 30

Table 11. Total sapling species abundance during two sampling periods ...... 44

Table 12. Small sapling species abundance during two sampling periods ...... 45

Table 13. Large sapling species abundance at two sampling periods ...... 46

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List of Figures

Figure 1. The Athens, Ironton, and Marietta Units of the Wayne National Forest (Hix and

Pearcy. 1997)...... 7

Figure 2. Diagram of an individual plot, comprised of the outer 500-m2 circular plot and inner 100-m2 circular plot (Hix and Chech, 1993) ...... 10

Figure 4. Relative densities of sapling species that were significantly different between

1993 and 2018 ...... 31

Figure 5. Stems per hectare of small-size sapling species that were significantly different between 1993 and 2018 ...... 32

Figure 6. Relative densities of small-size sapling species that were significantly different between 1993 and 2018 ...... 32

Figure 7. Stems per hectare of large-size sapling species that were significantly different between 1993 and 2018 ...... 33

Figure 8. Relative densities of large-size sapling species that were significantly different between 1993 and 2018 ...... 33

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Chapter 1. Introduction

1.1 Background

In recent decades, many eastern hardwood forests dominated by oak (Quercus) and hickory (Carya) species are not experiencing adequate regeneration of their overstory tree species. In contrast, mesophytic species including red maple (Acer rubrum), sugar maple (Acer saccharum), and American beech (Fagus grandifolia) are increasing in abundance in the regeneration layers (Nowacki and Abrams, 2008). These types of compositional changes are well documented in the following examples. In New York and

Massachusetts, red maple regeneration has increased in oak-dominated upland forests

(Lorimer, 1984). In dry-mesic ecosystems of the Wayne National Forest of southeastern

Ohio, very little oak sapling regeneration was found compared with the much more abundant saplings of several shade-tolerant genera (Acer and Fagus; Goebel and Hix,

1996). The present study found that while seedling-size regeneration of certain oak species were relatively common, there were few oak in the sapling layer.

This process of forest mesophication encompasses abiotic changes including increasing soil moisture, lower sunlight permeation into the understory, more fertile soils, and lower intensity of fires (Nowacki and Abrams, 2008). During the course of mesophication, the species richness of former oak-hickory dominated forest stands may initially increase with the developing regeneration of mesophytic species; eventually the 1 mesophytic species are expected to outcompete and succeed the established oak and hickory species (Abrams and Downs, 1990). A leading hypothesis regarding the cause of mesophication is alterations in fire regime (Abrams 1992, Lorimer 1993), although recent research has emphasized multiple factors including climate change (Delcourt et al. 1998), land-use changes (Dyer, 2010), and loss of keystone species (Ellison et al. 2005). The near extinction of the American chestnut (Castanea dentata) in the eastern United States created gaps suitable for oak: however, the mesophytic species sweet birch (Betula lenta),

Acer rubrum, and Fagus grandifolia have also benefited (Keever 1953, Woods 1959,

Good 1968). It can be argued that the Chestnut Blight (Cryphonectria parasitica) partially initiated the recent patterns in oak-maple dynamics (McEwan et al, 2011). In terms of changes in land use, oak have regenerated after human-orchestrated burns, with both

Native Americans (Gleason, 1913) and European settlers implementing fire regimes that were more severe and occurred at shorter intervals than those that previously had occurred naturally (Cole and Taylor, 1995). The eventual shift from these practices to fire suppression in the 1920s has been proposed as a leading contributor to the decline in oak, given its competitive adaptations to fire including thick bark, deep taproot, and high sprouting ability (Nowacki and Abrams, 2008). Mesophication hinders the more sun- loving or heliophytic species in the genera Quercus and Carya by theoretically operating as a positive feedback loop; mesophytic species which are favored by increasing shade, greater soil moisture and fertility, and low fire probability could also help increase these conditions through life-history traits including denser canopies and faster leaf litter composition (Alexander and Arthur, 2014).

Loss of oak in our forests is a concern for multiple reasons. Many wildlife species would lose an incredibly important resource, as acorns and leaves are widely consumed by eastern fauna (McShea et al. 2007). There is also a noteworthy human component to oak loss, as the timber is very valuable for use in barrel making, cabinetry, and flooring, among other uses (Shifley et al. 2006).

Mesophication remains a prominent lens for examining current shifts in eastern forest structure and species composition, and this has influenced our interest in changes in the mesophytic and heliophytic sapling species that are central to this paradigm.

. 1.2 Questions and Hypotheses

The overall goal of this thesis research is to examine the changes in the sapling layer of the Marietta Unit of the Wayne National Forest, in order to provide insight into the regeneration dynamics and future development of the forest ecosystems.

The research questions include:

1. Have there been shifts in sapling species abundance, from 1993 to 2018? Which species have experienced changes?

2. Are there changes in abundance of mesophytic sapling species, i.e., Acer rubrum, Acer saccharum, and Fagus grandifolia, and in heliophytic members of the genus Quercus?

The hypotheses are:

1.There have been statistically significant changes in sapling species composition and abundance between the two time periods; and

2. There have been significant changes in the regeneration of mesophytic species, i.e.,

Acer rubrum, Acer saccharum, and Fagus grandifolia, and also in Quercus spp.

Chapter 2. Study Area

The Wayne National Forest (WNF) in southeastern Ohio is comprised of the

Athens, Ironton, and Marietta Units (Figure 1). The study area is located in the part of the state known as the unglaciated Allegheny Plateau. This study occurred within the

Marietta Unit, the easternmost of the three units. The WNF is largely second-growth forest (Goebel and Hix, 1996) spanning 97,500 ha, with land in twelve counties. The area has a long history of resource extraction beginning with the settlement of Marietta in the late 1700s, which would eventually become the capital of the Northwest Territory

(Cayton, 1986). Much of the region was cleared by timber harvesting, agriculture, and mining of resources like iron and coal (Iverson et al, 2019). Many of these industries have since faltered, which lead to forest reestablishment in the late 1800s (Hutchinson et al, 2003).

2.1 Climate

The study area has a humid continental climate, which is characterized by warm summers and cold winters (National Oceanic and Atmospheric Administration, 2020).

Most precipitation falls in the form of rain averaging at 42.7 inches annually, and the 5 average temperature is 53.1 °F, with a winter low of 23.7 °F and summer high of 83.7 °F

(National Oceanic and Atmospheric Administration, 2020). Around half of the year, or

180 days, are frost free (Hix and Pearcy, 1997).

2.3 Geology and Soils

The bedrock of the Marietta Unit was formed during the Permian and

Pennsylvanian eras, dating back approximately 298-307 million years ago (Ohio Division of Geological Survey, 2017). The sedimentary bedrock is typically shale, sandstone, siltstone, mudstone, limestone, and coal (Lessig et al, 1977). Most of the local soils are comprised of loess, colluvium, and residuum of those bedrock types, and are typically 20-

40 inches deep (Hix and Pearcy, 1997). Soils range from well to moderately well-drained, and are classified as fine, fine loamy, mixed, mesic, typic Hapludalfs, and typic

Hapludalts (Hix and Pearcy, 1997).

2.4 Vegetation

The study area lies within the mixed-mesophytic forest region of Braun (1950).

The forests are primarily second-growth and of sprout origin after the widespread clearing of original mixed-oak, mixed-mesophytic, and beech stands by European immigrants in the late 1700s (Gordon, 1969). Tree species with the highest importance values are Acer rubrum and Quercus alba (Dyer, 2006).

Figure 1. The Athens, Ironton, and Marietta Units of the Wayne National Forest (Hix and Pearcy, 1997)

Chapter 3. Methods

3.1 Study Design

The Wayne National Forest Ecological Classification System (WNF ECS) project lead to the establishment of 370 permanent plots in the three forest administrative units during the years of 1993-1995 (Hix and Chech, 1993). The outcome of this project was to create classification systems for the ecosystem types of the Wayne National Forest. Field locations chosen for this study were second-growth mature stands, approximately a minimum of 70 years old when the permanent plots were established. Criteria for stand selection included: “(1) no obvious evidence of major anthropogenic disturbance within the past 40 years (e.g., no stumps), (2) no dominance of early successional tree species,

(3) no evidence of recent fire, significant windthrow, or unexplainable mortality, (4) stand is ‘fully stocked’ relative to the upland central hardwoods stocking chart (Gingrich,

1967), and (5) no extensive (multiple-tree) gaps in the canopy” (Hix and Pearcy, 1997).

Throughout the Marietta Unit, 128 permanent plots were established. The plots were located (often in pairs, but sometimes as many as clusters of five) along a total of 58 transects. With the use of GPS coordinates and plot descriptions, 98 plots were successfully relocated and resampled in the field seasons of 2017 and 2018 (Radcliffe, 8

2019). There were 30 plots that were harvested, or had since become inaccessible, and were not relocated.

3.2 Sampling Protocol

One aspect of the establishing and inventorying all ecosystem components of the permanent plots during the WNF ECS project of the 1990s, intensive sampling was conducted of all layers of the vegetation. In order to reference the plot center location, the nearest two or three ‘witness trees’ were noted, and a stake was placed to mark plot center. From that center, the radii of two concentric circles of 100 m2 within 500 m2 were measuring out by using Keson tapes (Figure 2). The 500-m2 plot is equivalent to 1/20th of a hectare, and 100-m2 plot is equivalent to 1/100th of a hectare. Saplings were recorded as those woody plant stems occurring within the 100-m2 inner circular plot. A sapling is defined as a woody plant taller than 1.37 m (breast height), with a DBH of 10.0 cm or less. Stems of this sapling size class were tallied by species. An additional size metric was distinguishing saplings up to 5.0 cm DBH as “small” and saplings between 5.1-10.0 cm as “large”. These field techniques and measurement implemented during in the first sampling period were replicated upon revisiting the plots to allow for direct comparisons between the two time periods.

Figure 2. Diagram of an individual plot, comprised of the outer 500-m2 circular plot and inner 100-m2 circular plot (Hix and Chech, 1993)

3.3 Data Analyses

The Wilcoxon signed-rank test was used to determine if the abundance of each sapling species significantly differed between the two sample years of 1993 and 2018.

This test is nonparametric, making it suitable for this dataset which is not normally distributed (Gibbons et al, 2003). A Wilcoxon signed-rank test is a paired difference test which assumes that the two samples are dependent. As the samples are the same 98 plots sampled at two different points in time, they are dependent variables. The Wilcoxon signed rank test was performed on each sapling species, using Minitab 17 software

(2010). No Bonferroni corrections or Bejamini-Hochberg procedures were performed in the analyses. In addition to testing the total counts for each species, the subcategories of small and large saplings were also tested, to see how the results may differ between size classes. The null hypothesis for each test is that the median difference between samples is zero, and the null alternative hypothesis is that the median difference between samples is not equal to zero. A significance level of P=0.05 was used. Therefore, if the P value was equal to or less than 0.05, the null hypothesis is rejected, and the null alternative is accepted.

Stems per hectare (SPH) was found by multiplying stem counts of total saplings

(Appendix A), small-size saplings (Appendix B), and large-size saplings (Appendix C) by an expansion factor which converts the species counts from the number of individuals of each species per study plot to number of individuals of each species per hectare

(United States Forest Service, 1965). We sampled 98 1/100th hectare plots, so the expansion factor 1.02 was used. Relative density (RD) for each species was found by dividing number of stems of a species by total number of stems in their sampling period and size class.

Chapter 4. Results

There were 46 species of saplings sampled during the two time periods (1993 and

2018) on the Marietta Unit of the Wayne National Forest (Table 1) Within the first sample, 42 species were found, and within the second sample, 40 species. The species

Castanea dentata, Carya laciniosa, Cercis canadensis, Kalmia latifolia, Morus rubra and

Pinus virginiana were unique to the 1993 sample, and the species Juglans cinerea,

Magnolia acuminata, Pinus strobus, and Viburnum dentatum were unique to the 2018 sample. In 1993, the three most common sapling species in descending order were Acer saccharum, Acer rubrum, and Cornus florida. Mesophytes Fagus grandifolia, Acer saccharum, and Acer rubrum were the top three most common sapling species sampled in

2018, also listed in descending order. Small-size saplings had the same order of rankings, with Acer saccharum, Acer rubrum, and Cornus florida being the most common in 1993. and Fagus grandifolia, Acer saccharum, and Acer rubrum being the most common species sampled in 2018 For the large-size class, Acer saccharum, Acer rubrum, and

Cornus florida were most common in 1993, and Acer saccharum, Acer rubrum, and

Fagus grandifolia were the most common in 2018.

Based on Wilcoxon signed-rank tests of the total numbers of saplings (of both size classes combined), there was a significant difference in each of the medians of seven 13 of the 46 present sapling species between the two time periods. The levels of significance by species were: Acer rubrum (P<0.001), Acer saccharum (P<0.001), Aesculus flava

(P=0.04), Carya glabra (P=0.01), Cornus florida (P<0.001), Fagus grandifolia (P=0.01), and Hamamelis virginiana (P=0.01)(Table 2).

Within the small sapling size-class, 10 species had significantly different median between the two time periods: Acer rubrum (P<0.001), Acer saccharum (P<0.001),

Carpinus caroliniana (P=0.05), Carya glabra (0.013), Cornus florida (P<0.001), Fagus grandifolia (P=0.05), Hamamelis virginiana (P=0.002), Nyssa sylvatica (P=0.01),

Prunus serotina (P=0.03), and Ulmus rubra (P=0.001) (Table 3).

There were only two species in the large sapling size-class with significantly different medians between the sample periods: Cornus florida (P=0.001) and Fagus grandifolia (P=0.002) (Table 4).

The total sapling count (Table 5) at the first sample in 1993 was 2706 SPH; there were only 1502 SPH during the second sample period in 2018. The species richness of the total sapling layer was 42 species in 1993. In 2018, the total sapling layer had 40 species. Fagus grandifolia increased from 225 to 307 SPH, and increased in RD from

8.31% to 20.44% (Table 6). Acer rubrum decreased from 459 SPH to 141 SPH and its relative density (RD) decreased from 16.96% to 9.39% in RD. Acer saccharum decreased from 603 to 267 SPH, and decreased from 22.28% to 17.78% in RD. Aesculus flava decreased from 16 to 5 SPH, and decreased from 0.59% to 0.33% RD. Carya glabra decreased from 40 SPH to 3, and decreased from 1.48% RD to 0.2%. Cornus florida decreased from 372 SPH to 6, and decreased from 13.75% to 0.4% in RD. Hamamelis

14 virginiana decreased from 114 to 61 SPH, and also decreased from 4.21% to 4.06% RD.

Comparison of SPH and RD values during both sampling periods for the seven species with statistically significant change are shown in Figures 3 and 4, respectively.

The number of small size-class saplings of all species in 1993 was 2452 SPH

(Table 7); small saplings decreased to 1278 SPH at the second sampling period in 2018.

Within the small size-class, there were 42 species present in the 1993 sample and 39 present in 2018. Fagus grandifolia increased from 206 to 259 SPH, and RD increased from 8.4% to 20.27% (Table 8). Acer rubrum decreased from 394 to 86 SPH, and decreased in RD from 16.06% to 6.7%. Acer saccharum decreased from 524 to 195 SPH, and decreased from 21.38% RD to 15.24%. Carpinus caroliniana decreased from 69 to

46 SPH, but increased from 2.83% to 3.59% RD. Carya glabra decreased from 36 to 2

SPH, and decreased from 1.46% to 0.16% RD. Cornus florida decreased from 345 to 4

SPH and also decreased from 14.06% to 0.32% RD. Hamamelis virginiana decreased from 111 to 60 SPH, though increased in RD from 4.53% to 4.71%. Nyssa sylvatica decreased from 51 to 14 SPH, and decreased in RD from 2.08% to 1.12%. Prunus serotina decreased from 15 to 4 SPH, and also decreased from 0.62% RD to 0.32%.

Ulmus rubra decreased from 83 to 6 SPH, and decreased from 3.37% to 0.48% RD.

Comparison of SPH and RD values during both sampling periods for the 10 small sapling species with statistically significant change are shown in Figures 5 and 6.

The number of large size-class saplings in 1993 was 254 SPH (Table 9), and it decreased to 224 SPH at the second sample in 2018. In this size class, there were 26 species present in 1993, and 21 present in 2018. Fagus grandifolia increased in SPH from

19 to 48, and increased in RD from 7.63% to 21.36% (Table 10). Cornus florida decreased in SPH from 28 to 2, and RD decreased from 10.84% to 0.91%. Comparison of

SPH and RD values during both sampling periods for the two large sapling species with statistically significant change are shown in Figures 7 and 8.

No species of Quercus was found to have had significant change at the total, small-, or large-size classes. This contrasts with mesophytic species Fagus grandifolia, which differed in the total, small, and large categories, as well as Acer rubrum and Acer saccharum which both differed in the total and small size categories.

Chapter 5. Discussion

There were decreasing abundances and relative densities of total, small- and large-size saplings of Acer rubrum. Acer saccharum decreased in abundance and relative density in the total and small-size classes. In the large-size class, relative density of Acer saccharum increased, with decrease in abundance. Lindera benzoin saplings were more numerous and had higher relative density values than Acer rubrum in total and small size classes; however, Lindera benzoin is a shrub species. The changes in the large-size classes of Acer rubrum and Acer saccharum were not statistically significant.

. In contrast, both relative densities and stem counts significantly increased for all three size classes of Fagus grandifolia. These three mesic species of Fagus grandifolia, Acer saccharum, and Acer rubrum were the highest in abundance and relative density among tree species in 2018.

The trend of increases in Fagus abundances relative to Acer is not generally associated with mesophication in the central hardwood region (Nowacki and Abrams,

2008). However, other studies in southeastern Ohio have also observed similar trends. On both the Athens Unit of the Wayne National Forest (Palus et al. 2018), and in unburned stands of Vinton Furnace State Experimental Forest (Hutchinson et al. 2012), Fagus grandifolia was the most common sapling species. The change in Fagus regeneration 17 may be due to the root suckers of this species increasing their competitiveness relative to

Acer, as has been observed in other central hardwoods forests including minimally disturbed stands in northern Alabama (Richards and Hart, 2011). On the Athens Unit of the Wayne, it was hypothesized that factors like fire suppression and Acer rubrum abundance ultimately help create more mesic conditions that are more suited for Fagus regeneration (Palus et al. 2018). Given the observed high survival rates of sapling Fagus grandifolia and the potential for ingrowth (Gauthier et al. 2015, Burns and Honkala,

1990), it is possible the future canopy tree species composition on the Marietta Unit will include higher dominance by Fagus grandifolia.

Fagus grandifolia may be impacted in the future by beech-bark disease (Cale et al. 2017) or beech leaf disease (Ewing et al. 2018), both of which could impact the survival rates of the species in Ohio. It is worth noting that in the other recent studies that have also noted increases in beech regeneration in the northeastern hardwood forests of

Quebec (Duchesne and Ouimet, 2009, Gravel et al, 2011), increased sprouting is a response associated with beech bark disease damage (Hane, 2004). It remains to be seen how these pathogens and other threats may affect the probabilities of saplings of Fagus grandifolia and other mesophytic species ascending into the canopy.

Examining the trends in the regeneration of Quercus saplings remains complicated given their small sample sizes in this study. Our analyses did not find the difference in medians of any Quercus spp. to be statistically significant. It is important to note that the number of Quercus stems appear to be overall decreasing over time on the

Marietta Unit, although it can be difficult to detect trends given the small sample sizes.

Some species did see increases in SPH or RD, like Quercus rubra, but these occur on very small scales. For example, large Quercus rubra saplings increased from 2 to 4 SPH and 0.8% to 1.82% RD from 1993 to 2018. Of our hickory species and heliophytic associates, Carya glabra did show significant changes in the total sapling and small sapling categories, where SPH and RD decreased in both total and small classes.

Cornus florida experienced the most extreme decline in this study, both in number of stems and relative density. This change is likely due to the dogwood anthracnose (Discula destructiva).The once abundant tree species has become less common, as this lethal disease has killed as many as 90% of Cornus in certain forest stands (Holzmueller et al, 2006). Similar decreases in this species were observed in the

Athens Unit of the WNF (Palus et al. 2018). A study in southern Illinois spanning 1994 to

2004 (Sucheki and Gibson, 2008) linked dogwood loss with increased densities in other species (Asimina triloba, Sassafras albidum, Acer rubrum, and Ostrya virginiana). Their reported trends are not the same as reported here, although Sassafras albidum and Acer rubrum did increase in abundance on the Marietta Unit.

Species which were present in one sampling period and not the other were examined for possible explanations. Unfortunately, the species that did not occur during both time periods had very small samples sizes that made their trends difficult to interpret. One consideration was if the species was known to be uncommon and (or) a species of concern. For example, Carya laciniosa, Cercis canadensis, Kalmia latifolia,

Magnolia acuminata, Morus rubra, Pinus virginiana, Pinus strobus, and Viburnum dentatum were each only found in one sample period. However, these species are all

19 considered secure (International Union for Conservation of Nature 2020); therefore, presence or absence may not be an indicator, especially when there have been as few as one stem sampled in some cases.

Castanea dentata and Juglans cinerea (butternut) were more closely examined, as the species both suffer from respective diseases (Schlarbaum et al. 1997). Castanea dentata, although critically endangered (International Union for Conservation of Nature,

2020), can be found as sprout-origin stems sporadically across its native range. Juglans cinerea appeared even less frequently than American chestnut in the study plots. This species is currently endangered (International Union for Conservation of Nature, 2020) due to the fungal butternut canker disease (Sirococcus clavigignenti-juglandacearum).

Again, while the sample sizes are not large enough to draw conclusions, it is of interest to have knowledge of the whereabouts of threatened species, as this information may be able to contribute to efforts to protect and restore these species.

Our study sought determine if changes have occurred in these stands between study periods. By testing for population differences, and quantitatively evaluating them at the species level, we were able to explore shifts in sapling abundance and composition on the Marietta Unit of the Wayne National Forest.

Table 1. Species of sapling size sampled during the two time periods (1993 and 2018) on the Marietta Unit of the Wayne National Forest by Latin name and species code

Latin name Species code Acer negundo acne Acer rubrum acru Acer saccharum acsa Aesculus flava aefl Aesculus glabra aegl Amelanchier arborea amar Asimina triloba astr Carpinus caroliniana caca Carya cordiformis caco Castanea dentata cade Carya glabra cagl Carya laciniosa cala Carya ovata caov Carya tomentosa cato Cercis canadensis ceca Corylus americana coam Cornus florida cofl Crataegus spp. cr spp Fagus grandifolia fagr Fraxinus americana fram Fraxinus pennsylvanica frpe Hamamelis virginiana havi Kalmia latifolia kala Juglans cinerea juci Lindera benzoin libe Liriodendron tulipifera litu Magnolia acuminata maac Morus rubra moru Nyssa sylvatica nysy Ostrya virginiana osvi Oxydendrum arboreum oxar Pinus strobus pist Pinus virginiana pivi Prunus serotina prse Quercus alba qual Quercus montana qumo Quercus rubra quru Quercus velutina quve Sassafras albidum saal Tilia americana tiam Tsuga canadensis tcsa Ulmus americana ulam Ulmus rubra ulru Viburnum acerifolium viac Viburnum dentatum vide Viburnum prunifolium vipr

Table 2. Wilcoxon signed-rank test results for all saplings (both size classes combined) Species code P-Value acne 0.37 acru <0.001 acsa <0.001 aefl 0.04 aegl 0.59 amar 0.43 astr 0.67 caca 0.12 caco 0.4 cade 0.18 cagl 0.01 cala 1 caov 1 cato 1 ceca 1 coam 0.37 cofl <0.001 cr spp 0.59 fagr 0.01 fram 0.33 frpe 0.15 havi 0.01 kala 1 juci 1 libe 0.4 litu 0.04 maac 1 moru 1 nysy 0.02 osvi 0.07 oxar 0.36 pist 1 pivi 0.37 prse 0.03 qual 0.4 qumo 0.83 quru 0.21 quve 0.11 saal 0.97 tiam 0.23 tcsa 0.37 ulam 0.46 ulru 0.001 viac 0.22 vide 1 vipr 1

Table 3. Wilcoxon signed-rank test results for small saplings Species code P-Value acne 0.37 acru <0.001 acsa <0.001 aefl 0.06 aegl 0.86 amar 0.43 astr 0.87 caca 0.05 caco 0.4 cade 0.18 cagl 0.013 cala 1 caov 0.79 cato 0.45 ceca 1 coam 0.37 cofl <0.001 cr spp 0.79 fagr 0.05 fram 0.47 frpe 0.31 havi 0.002 kala 1 juci 1 libe 0.41 litu 0.06 maac 1 moru 1 nysy 0.01 osvi 0.18 oxar 0.86 pist 1 pivi 0.37 prse 0.03 qual 0.86 qumo 0.94 quru 0.17 quve 0.18 saal 0.76 tiam 0.47 tcsa 0.37 ulam 1 ulru 0.001 viac 0.13 vide 1 vipr 1

Table 4. Wilcoxon signed-rank test results for large saplings Species code P-Value acne 1 acru 0.31 acsa 0.64 aefl 0.37 aegl 0.37 amar 1 astr 1 caca 0.83 caco 1 cade 1 cagl 0.28 cala 1 caov 1 cato 0.36 ceca 1 coam 1 cofl 0.001 cr spp 1 fagr 0.002 fram 0.37 frpe 0.18 havi 0.37 kala 1 juci 1 libe 1 litu 0.59 maac 1 moru 1 nysy 0.37 osvi 0.35 oxar 1 pist 1 pivi 0.37 prse 1 qual 0.36 qumo 1 quru 0.85 quve 0.37 saal 0.37 tiam 0.37 tcsa 1 ulam 0.5 ulru 0.37 viac 1 vide 1 vipr 0.181

Table 5. Sapling (both size classes combined) stems per hectare (SPH) by year of sampling

Species code 1993 2018 acne 3 1 acru 459 141 acsa 603 267 aefl 16 5 aegl 5 12 amar 20 13 astr 94 66 caca 73 49 caco 3 6 cade 3 0 cagl 40 3 cala 2 0 caov 3 3 cato 15 15 ceca 1 0 coam 7 21 cofl 372 6 cr spp 7 4 fagr 225 307 fram 33 37 frpe 16 43 havi 114 61 kala 2 0 juci 0 1 libe 181 234 litu 24 11 maac 0 1 moru 1 0 nysy 52 18 osvi 91 46 oxar 14 21 pist 0 3 pivi 10 0 prse 15 4 qual 12 8 qumo 13 6 quru 31 18 quve 18 6 saal 9 15 tiam 8 4 tcsa 5 1 ulam 3 9 ulru 91 6 viac 5 15 vide 0 4 vipr 3 7 Total 2706 1502

Table 6. Relative density % (RD) of species for all saplings (both size classes combined)

Species code 1993 total 2018 total acne 0.11 0.07 acru 16.96 9.39 acsa 22.28 17.78 aefl 0.59 0.33 aegl 0.18 0.8 amar 0.74 0.87 astr 3.47 4.39 caca 2.7 3.26 caco 0.11 0.4 cade 0.11 0 cagl 1.48 0.2 cala 0.07 0 caov 0.11 0.2 cato 0.55 1 ceca 0.04 0 coam 0.26 1.4 cofl 13.75 0.4 cr spp 0.26 0.27 fagr 8.31 20.44 fram 1.22 2.46 frpe 0.59 2.86 havi 4.21 4.06 kala 0.07 0 juci 0 0.07 libe 6.69 15.58 litu 0.89 0.73 maac 0 0.07 moru 0.04 0 nysy 1.92 1.2 osvi 3.36 3.06 oxar 0.52 1.4 pist 0 0.2 pivi 0.37 0 prse 0.55 0.27 qual 0.44 0.53 qumo 0.48 0.4 quru 1.15 1.2 quve 0.67 0.4 saal 0.33 1 tiam 0.3 0.27 tcsa 0.18 0.07 ulam 0.11 0.6 ulru 3.36 0.4 viac 0.18 1 vide 0 0.27 vipr 0.11 0.47

Table 7. Small saplings per hectare (SPH) by year of sampling

Species code 1993 2018 acne 2 0 acru 394 86 acsa 524 195 aefl 12 3 aegl 4 9 amar 20 13 astr 94 65 caca 69 46 caco 3 6 cade 3 0 cagl 36 2 cala 2 0 caov 3 2 cato 14 9 ceca 1 0 coam 6 21 cofl 345 4 cr spp 6 4 fagr 206 259 fram 31 37 frpe 16 40 havi 111 60 kala 2 0 juci 0 1 libe 181 234 litu 21 10 maac 0 1 moru 1 0 nysy 51 14 osvi 81 43 oxar 12 16 pist 0 3 pivi 10 0 prse 15 4 qual 7 6 qumo 12 6 quru 29 14 quve 16 6 saal 7 15 tiam 6 4 tcsa 5 1 ulam 1 4 ulru 83 6 viac 5 15 vide 0 4 vipr 3 7 Total 2452 1278

Table 8. Relative density % (RD) of species for small saplings by year of sampling

Species code 1993 2018 acne 0.08 0 acru 16.06 6.7 acsa 21.38 15.24 aefl 0.5 0.24 aegl 0.17 0.72 amar 0.83 1.04 astr 3.83 5.11 caca 2.83 3.59 caco 0.12 0.48 cade 0.12 0 cagl 1.46 0.16 cala 0.08 0 caov 0.12 0.16 cato 0.58 0.72 ceca 0.04 0 coam 0.25 1.68 cofl 14.06 0.32 cr spp 0.25 0.32 fagr 8.4 20.27 fram 1.25 2.87 frpe 0.67 3.11 havi 4.53 4.71 kala 0.08 0 juci 0 0.08 libe 7.36 18.28 litu 0.87 0.8 maac 0 0.08 moru 0.04 0 nysy 2.08 1.12 osvi 3.29 3.35 oxar 0.5 1.28 pist 0 0.24 pivi 0.42 0 prse 0.62 0.32 qual 0.29 0.48 qumo 0.5 0.48 quru 1.16 1.12 quve 0.67 0.48 saal 0.29 1.2 tiam 0.25 0.32 tcsa 0.21 0.08 ulam 0.04 0.32 ulru 3.37 0.48 viac 0.21 1.2 vide 0 0.32 vipr 0.12 0.56

Table 9. Large saplings per hectare (SPH) by year of sampling

Species code 1993 2018 acne 1 1 acru 65 55 acsa 79 72 aefl 4 2 aegl 1 3 amar 0 0 astr 0 1 caca 4 3 caco 0 0 cade 0 0 cagl 4 1 cala 0 0 caov 0 1 cato 1 6 ceca 0 0 coam 1 0 cofl 28 2 cr spp 1 0 fagr 19 48 fram 2 0 frpe 0 3 havi 3 1 kala 0 0 juci 0 0 libe 0 0 litu 3 1 maac 0 0 moru 0 0 nysy 1 4 osvi 10 3 oxar 2 5 pist 0 0 pivi 0 0 prse 0 0 qual 5 2 qumo 1 0 quru 2 4 quve 2 0 saal 2 0 tiam 2 0 tcsa 0 0 ulam 2 5 ulru 8 0 viac 0 0 vide 0 0 vipr 0 0 Total 254 224

Table 10. Relative density % (RD) of species for large saplings by year of sampling

Species code 1993 2018l acne 0.4 0.45 acru 25.7 24.55 acsa 30.92 32.27 aefl 1.61 0.91 aegl 0.4 1.36 amar 0 0 astr 0 0.45 caca 1.61 1.36 caco 0 0 cade 0 0 cagl 1.61 0.45 cala 0 0 caov 0 0.45 cato 0.4 2.73 ceca 0 0 coam 0.4 0 cofl 10.84 0.91 cr spp 0.4 0 fagr 7.63 21.36 fram 0.8 0 frpe 0 1.36 havi 1.2 0.45 kala 0 0 juci 0 0 libe 0 0 litu 1.2 0.45 maac 0 0 moru 0 0 nysy 0.4 1.82 osvi 4.02 1.36 oxar 0.8 2.27 pist 0 0 pivi 0 0 prse 0 0 qual 2.01 0.91 qumo 0.4 0 quru 0.8 1.82 quve 0.8 0 saal 0.8 0 tiam 0.8 0 tcsa 0 0 ulam 0.8 2.27 ulru 3.21 0 viac 0 0 vide 0 0 vipr 0 0

Stems Per Hectare of Total Saplings 700

600

500

400

300

200 Stems per Hectare

100

0 Acer rubrum Acer Aesculus flava Carya glabra Cornus florida Fagus Hamamelis saccharum grandifolia virginiana

Sapling Species 1993 2018

Figure 3. Total stems per hectare of sapling species that were significantly different between 1993 and 2018

Relative Density of Total Saplings 25.00%

20.00%

15.00%

10.00%

5.00% Percent Relative Density

0.00% Acer rubrum Acer Aesculus Carya glabra Cornus Fagus Hamamelis saccharum flava florida grandifolia virginiana

Sapling Species 1993 2018

Figure 4. Relative densities of sapling species that were significantly different between 1993 and 2018

Stems per Hectare of Small Saplings 600 500 400 300 200

Stems per Hectare 100 0

Acer rubrum Ulmus rubra Carya glabraCornus florida Acer saccharum Nyssa sylvaticaPrunus serotina Fagus grandifolia Carpinus caroliniana Hamamelis virginiana

Sapling Species 1993 2018

Figure 5. Stems per hectare of small-size sapling species that were significantly different between 1993 and 2018

Relative Density of Small Saplings 25.00%

20.00%

15.00%

10.00%

5.00%

Percent Relative Density 0.00%

Acer rubrum Ulmus rubra Carya glabraCornus florida Acer saccharum Nyssa sylvaticaPrunus serotina Fagus grandifolia Carpinus caroliniana Hamamelis virginiana

Sapling Species 1993 2018

Figure 6. Relative densities of small-size sapling species that were significantly different between 1993 and 2018

Stems per Hectare of Large Saplings 30

10 Stems per Hectare

0 Cornus florida Fagus grandifolia

Sapling Species 1993 2018

Figure 7. Stems per hectare of large-size sapling species that were significantly different between 1993 and 2018

Relative Density of Large Saplings 25.00%

20.00%

15.00%

10.00%

Relative Density Percent 5.00%

0.00% Cornus florida Fagus grandifolia

Sapling Species 1993 2018

Figure 8. Relative densities of large-size sapling species that were significantly different between 1993 and 2018

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Appendix: Sapling species abundance during two sampling periods

Table 11. Total sapling species abundance during two sampling periods

Species code 1993 2018 acne 3 1 acru 450 138 acsa 591 262 aefl 16 5 aegl 5 12 amar 20 13 astr 92 65 caca 72 48 caco 3 6 cade 3 0 cagl 39 3 cala 2 0 caov 3 3 cato 15 15 ceca 1 0 coam 7 21 cofl 365 6 cr spp 7 4 fagr 221 301 fram 32 36 frpe 16 42 havi 112 60 kala 2 0 juci 0 1 libe 177 229 litu 24 11 maac 0 1 moru 1 0 nysy 51 18 osvi 89 45 oxar 14 21 pist 0 3 pivi 10 0 prse 15 4 qual 12 8 qumo 13 6 quru 30 18 quve 18 6 saal 9 15 tiam 8 4 tcsa 5 1 ulam 3 9 ulru 89 6 viac 5 15 vide 0 4 vipr 3 7 total 2653 1473

Table 12. Small sapling species abundance during two sampling periods

Species code 1993l 2018 acne 2 0 acru 386 84 acsa 514 191 aefl 12 3 aegl 4 9 amar 20 13 astr 92 64 caca 68 45 caco 3 6 cade 3 0 cagl 35 2 cala 2 0 caov 3 2 cato 14 9 ceca 1 0 coam 6 21 cofl 338 4 cr spp 6 4 fagr 202 254 fram 30 36 frpe 16 39 havi 109 59 kala 2 0 juci 0 1 libe 177 229 litu 21 10 maac 0 1 moru 1 0 nysy 50 14 osvi 79 42 oxar 12 16 pist 0 3 pivi 10 0 prse 15 4 qual 7 6 qumo 12 6 quru 28 14 quve 16 6 saal 7 15 tiam 6 4 tcsa 5 1 ulam 1 4 ulru 81 6 viac 5 15 vide 0 4 vipr 3 7 total 2404 1253

Table 13. Large sapling species abundance at two sampling periods

Species code 1993 2018 acne 1 1 acru 64 54 acsa 77 71 aefl 4 2 aegl 1 3 amar 0 0 astr 0 1 caca 4 3 caco 0 0 cade 0 0 cagl 4 1 cala 0 0 caov 0 1 cato 1 6 ceca 0 0 coam 1 0 cofl 27 2 cr spp 1 0 fagr 19 47 fram 2 0 frpe 0 3 havi 3 1 kala 0 0 juci 0 0 libe 0 0 litu 3 1 maac 0 0 moru 0 0 nysy 1 4 osvi 10 3 oxar 2 5 pist 0 0 pivi 0 0 prse 0 0 qual 5 2 qumo 1 0 quru 2 4 quve 2 0 saal 2 0 tiam 2 0 tcsa 0 0 ulam 2 5 ulru 8 0 viac 0 0 vide 0 0 vipr 0 0 total 249 220